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Context-Dependent Effects of Nutrition and Dam Behavior on Neonatal Survival in a Long-Lived Herbivore

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Context-Dependent Effects of Nutrition and Dam Behavior on Neonatal Survival in a Long-Lived Herbivore Context-Dependent Effects of Nutrition and Dam Behavior on Neonatal Survival in a Long-Lived Herbivore A Thesis Presented in Partial Fulfillment of the Requirements for the Degree of Master of Science with a Major in Natural Resources in the College of Graduate Studies University of Idaho by Nicole M. Bilodeau Major Professor: Ryan A. Long, Ph.D. Committee Members: E. Frances Cassirer, Ph.D.; Sophie L. Gilbert, Ph.D.; Lisa A. Shipley, Ph.D. Department Administrator: Lisette P. Waits, Ph.D. August 2021 ii Authorization to Submit Thesis This thesis of Nicole M. Bilodeau, submitted for the degree of Master of Science with a Major in Natural Resources and titled “Context-Dependent Effects of Nutrition and Dam Behavior on Neonatal Survival in a Long-Lived Herbivore,” has been reviewed in final form. Permission, as indicated by the signatures and dates below, is now granted to submit final copies to the College of Graduate Studies for approval. Major Professor: _____________________________________ Date: __________ Ryan A. Long, Ph.D. Committee Members: _____________________________________ Date: __________ E. Frances Cassirer, Ph.D. _____________________________________ Date: __________ Sophie L. Gilbert, Ph.D. _____________________________________ Date: __________ Lisa A. Shipley, Ph.D. Department Administrator: _____________________________________ Date: __________ Lisette P. Waits, Ph.D. iii Abstract Behavior represents one of the primary mechanisms by which animals overcome environmental constraints on survival and reproductive success. Females in particular often exhibit plastic behavioral strategies for coping with the different nutritional demands and degrees of susceptibility to predation imposed by gestation, parturition and lactation. Previous studies have demonstrated a link between space-use behavior and important correlates of fitness and have highlighted the value of mechanistic nutritional approaches for understanding the fitness consequences of behavior. However, the mechanisms by which individual responses to variation in the nutritional landscape scale up to influence population performance remain unclear. We quantified relationships among the nutritional landscape (i.e., spatiotemporal variation in forage biomass), dam behavior, and neonatal survival in bighorn sheep (Ovis canadensis). We conducted intensive vegetation sampling and used generalized additive modeling to map the nutritional landscapes available to sheep during summer (May–September) in three population ranges in Idaho: Owyhee River, East Fork of the Salmon River, and Lost River Range. We used GPS collars and lamb surveys to monitor ewe behavior and lamb survival in each study area, and used known-fate survival modeling to test for behaviorally mediated effects of nutrition on lamb survival. Relationships among the nutritional landscape, ewe behavior, and lamb survival were context dependent and varied among study sites. In the Lost River, where lamb survival was highest (83.9%), probability of lamb survival increased when ewes traded access to rugged terrain for access to higher forage biomass. We observed the opposite pattern in the East Fork (i.e., probability of lamb survival increased when ewes traded access to forage for access to rugged terrain), however, and in the Owyhee no metric of ewe behavior was significantly related to the probability of lamb survival. We also observed a strong, positive relationship between spring nutritional condition and probability of lamb survival across study sites. Our research helps to establish mechanistic links among habitat heterogeneity, individual space-use behavior, and reproductive success in bighorn sheep, and underscores the fundamental importance of nutrition as a driver of ungulate performance. Continuing to improve our understanding of such relationships will provide valuable insights for managers and conservationists, and will aid in accurately parameterizing models of population dynamics. Maximizing the usefulness of such models requires knowledge of the mechanisms that underpin variation in population iv demographics, and nutritional-ecological approaches like those used in our study shed important light on those mechanisms. v Acknowledgments The number of people to whom I owe the success of this project could be a chapter in and of itself. First, the support and collaboration between the University of Idaho and the Idaho Department of Fish and Game was instrumental in the success of this study. I was exceptionally fortunate to have Ryan Long as my major professor who bolstered my moral during times of frustration, expanded my critical thinking into the bigger picture, and inspired me with his passion for wildlife science. I would like to express my deepest appreciation to my committee members Frances Cassirer, Sophie Gilbert, Lisa Shipley, and Ryan Long for their exceptional expertise, their valuable project recommendations, and their constructive edits throughout the writing process. I would also like to extend my deepest gratitude to Mark Hurley who saw potential in me and took the time to mentor me over the years, which is how I got to where I am today. It was a privilege to be a part of the “Hurley Circus” and I learned more than I could have ever imagined including how valuable my weight is in jet fuel. I am extremely grateful to the entire bighorn sheep capture crew including but not limited to Paul Atwood, Curtis Hendricks, Bret Panting, Shane Roberts, Stacey Dauwalter, John Nelson, Josh Rydalch, Brett Stansberry, Jessie Shallow, Chris Gaughn, Chel Curtis, Katie Oelrich, Michelle Kemner, Brian Marek, Trent Brown, Nathan Borg, Sierra Robatcek, and Cindy McClellan. Your immense hard work, enthusiastic comradery, and endless humor made the 16 hour days just fly by! A special thanks to Hollie Miyasaki for being an excellent capture coordinator and providing a wealth of data. I must also thank Dr. Mark Drew for his uncompromising care for wildlife and for taking the time to teach. I’d also like to extend my gratitude to Tom Stephenson for assisting with capture, providing ultrasound training, and imparting his wisdom of all things sheep. I am deeply grateful to our helicopter pilot Tony Herby who not only made sure we all made it home safe every night, but also provided endless entertainment while on the road. I would like to extend my sincere thanks to Chel Curtis and Brett Stansberry for their invaluable insight and contribution to my project planning, logistics, sheep capture, and surveys. I must also thank Greg Painter, Dennis Newman, Michelle Kemner, and Curtis Hendricks for their help coordinating regional staff and assisting with lamb surveys. A special thank you to John Romero at Owyhee Air for being a smooth pilot, a great lamb spotter, and for bringing me breakfast cookies! Many thanks to two of the greatest botany extraordinaires, Lynn Kinter and Chris Murphy, for all of vi your valuable insight, field training, and for inspiring me to love plants. A big thank you to the staff at the Wildlife Health Lab including but not limited to Stacey Dauwalter, Tricia Hebdon, and Shayla March for processing plant samples, letting me store endless amounts of equipment at the lab, making room for my mobile office, and for being awesome people to work with. I am deeply indebted to Doug Engemann and Morgan Fife at the Pahsimeroi Fish Hatchery for providing free housing, sample storage, and the perfect staging area for my summer field work. They were always welcoming and eager to help with inevitable equipment failures, and for that I will always be grateful! I owe a thank you to the Baker family and the Payne family for granting us permission onto your property to access bighorn sheep herds that would otherwise be nearly impossible to track. I would also like to thank Tony Folsom at the Clearwater Fish Hatchery for sharing the ever coveted freezer space with my hundreds of plant samples. Thank you to Jon Horne for letting me borrow your truck for unexpected and last minute transport trips as well as hosting great BBQs and providing invaluable R code assistance. I want to thank Dorah Mtui for going out of her way to help coordinate equipment storage space and laboratory access for me. My deepest appreciation goes to Lisa Shipley for her indispensable expertise and for teaching me how to run nutritional assays at WSU. I am particularly grateful to John Fluegel at the Steffen Center for tolerating the sound of a plant grinder for hundreds of hours and always letting me stay late. I also appreciate all the volunteer coordination and advice I received from Bill London and Zach Higgins at the Wild Sheep Foundation. I owe a big thank you to the countless volunteers who helped with lamb surveys all across the state, and while I didn’t get to meet all of you this project would not have been possible without you. In addition to all the volunteers, there were many IDFG staff members who spent long days hiking in rough sheep country to help complete surveys including Sarah Meronk, Ian Montgomery, Cathy Henry, Bret Stansberry, John Nelson, Morgan Pfander, Iver Hull, Brendon Oats, Paul Page, Dane Cook, Dustin Brewster, Tyler Rothe, and many more! I cannot begin to express my thanks to my technicians Andrea Gibbons, Liana Cabiles, Brendon Harker, Colby Slezak, Andy Landsem, Emily Holmes, Layne Saidnawey, and Conway LeBleu for your dedication and hard work in the field. I promise we really needed all those plants I made you pick! A special thanks to Matt Rafferty for working with me during the winter processing
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